We consider 27 population and community terms used frequently by parasitologists when describing the ecology of parasites. We provide suggestions for various terms in an attempt to foster consistent use and to make terms used in parasite ecology easier to interpret for those who study free-living organisms. We suggest strongly that authors, whether they agree or disagree with us, provide complete and unambiguous definitions for all parameters of their studies.

SUMMARYRecently, some authors (Kennedy, 1981; Price & Clancy, 1983) have argued that there are fundamental differences between the communities of helminths in fish and bird hosts. Such differences are foreshadowed by the work of Dogiel (1964) and are apparent from survey data (e.g. Threlfall, 1967; Bakke, 1972; Hair & Holmes, 1975 on birds, and compare Chubb, 1963; Mishra & Chubb, 1969; Wootten, 1973; Ingham & Dronen, 1980 on fish). Questions still remain, however, as to whether the distinctions are truly justified and whether the differences are really fundamental. In this paper, we address these questions by examining helminth diversity in a series of hosts. We then discuss and provide explanations for the observed differences.

SummaryWe examine patterns of community richness among intestinal parasitic helminth communities in fishes, herptiles, birds and mammals with respect to the comparative number of component species in a host population. We show that terrestrial hosts have, on average, fewer component species than aquatic hosts. We also show that the mean number of component species in aquatic hosts increases from fishes through herptiles to birds before declining slightly in mammals. For terrestrial hosts, the mean number of component species increases from herptiles, through birds, reaching a maximum in mammals. We conclude that: (i) habitat of the host is more important in determining community richness than is host phylogeny; (ii) the phenomenon of 'host capture' may be largely responsible for increased species richness in some host groups; (iii) aquatic birds harbour the richest intestinal helminth communities; and (iv) as we interpret them, our data refute the time hypothesis, which would predict that fishes as the oldest lifestyle should have the richest helminth communities.

Examples of the apparently stochastic nature of freshwater fish helminth communities illustrating the erratic and unpredictable occurrence and distribution of many species are provided for six species of fish from several localities throughout Britain. By focussing on parasite colonization strategies two categories of helminths are recognized: autogenic species which mature in fish and allogenic species which mature in vertebrates other than fish and have a greater colonization potential and ability. Three groups of fish are distinguished: salmonids, in which helminth communities are generally dominated by autogenic species which are also responsible for most of the similarity within and between localities; cyprinids, in which they are dominated by allogenic species which are also responsible for most of the similarity within and between localities; and anguillids, whose helminth communities exhibit intermediate features with neither category consistently dominating nor providing a clear pattern of similarity. Recognition and appreciation of the different colonization strategies of autogenic and allogenic helminths in respect of host vagility and ability to cross land or sea barriers and break down habitat isolation, and their period of residence in a locality, whether transient or permanent, provides an understanding of, and explanation for, the observed patchy spatial distribution of many helminths. Comparison with other parts of the world indicates that colonization is a major determinant of helminth community structure.

This paper examines patterns in the linear distribution of helminths in the small intestine of each of 45 scaup. Over all birds, most common species of helminths were found in over half of the small intestine; their distributions overlapped broadly. In individual birds, each species occupied a more restricted, predictable part of the intestine; "core species" (those present in more than two-thirds of the birds) were more evenly distributed along the intestine than expected by chance and occupied almost all of it; "secondary species" (those in one- to two-thirds of the birds) were clumped in anterior and posterior portions of the intestine; and "satellite species" (those in less than one-third of the birds) were randomly distributed. For all common helminths, the range occupied in individual birds was significantly correlated with population size; however, overlap between adjacent species did not increase with increases in their total numbers. The intestinal helminths of scaup belonged to three guilds: small absorbers (paramucosal), large absorbers (mid-lumenal), and trematodes. The interactions noted above occurred both within and between guilds. The overall helminth community in scaup appeared to be saturated with species belonging to the two absorber guilds, but not with trematodes. Communities within individual scaup sampled this overall community and were often unsaturated. We conclude that the core and secondary species of absorbers provide a basic, interactive structure to the overall community and to most of the communities in individual birds. Trematodes and satellite absorber species provide stochastic elaborations.

The helminth communities of the small intestines of 45 lesser scaup ducks sampled from 13 lakes in Alberta, Canada, were examined for patterns of association among species. A core of eight frequent, numerous, and positively associated species provided a basic similarity across all host individuals. Six of these core species are specialists in lesser scaup, and the other two are generalists in waterfowl. A group of eight moderately frequent and numerous species, which were positively associated with the core species but not with each other, also contributed to this similarity; these secondary species included two specialists in scaup and three generalists in waterfowl. The 36 remaining "satellite species" appeared to be distributed randomly among birds. Variations among communities in individual birds were due largely to (i) differences in the numbers of two suites of helminth species, one using Hyalella azteca and the other Gammarus lacustris as intermediate host; (ii) differences in the numbers of the Hyalella suite, and in the presence or absence of some secondary or satellite species, associated with the lake from which the duck was taken; and (iii) differences in the total numbers of helminths per duck, which may be associated with differential susceptibility of individual ducks.

Problems of pattern and scale are considered in relation to helminth communities of freshwater fish by examining them at different hierarchical taxonomic and spatial scales, with a view to seeking generalizations of heuristic value, assessing the importance of phylogenetic and ecological determinants of community structure and improving understanding of unpredictable communities. Initially, communities were analysed at the level of salmonid genera, focusing on Oncorhynchus, in its heartland in Canada: then in O. mykiss throughout its global range and finally in individual localities to which it has been introduced in Britain. In the heartland, communities are dominated by salmonid specialist helminths, forming a phylogenetic element: the minority ecological element comprises broad generalists and non-salmonid specialists. Most species except generic specialists are shared between host genera. As the distance to which O. mykiss was translocated from its heartland increases, so generic specialists disappear first and then salmonid specialists decline. The community is thus increasingly composed of generalists and it also becomes increasingly poor. Helminths may be acquired from native salmonids and/or unrelated hosts, depending on availability. This same pattern is paralleled in individual localities in a restricted region: the phylogenetic element reflects the native salmonid species present and the ecological element the presence of other genera of fish; i.e., a supply-side situation. The change of scale in analysis has thus enabled the recognition of generalizations and patterns of heuristic value and improved the understanding of unpredictable communities by interpreting local variation as ecological 'noise' that often obscures fundamental patterns. In this and other taxa of fish, phylogenetic elements dominate helminth communities in the heartlands, but ecological elements dominate as the host increasingly becomes a stranger in a strange land.

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